All sequencing data produced in the present study have been deposited in the NCBI Short Read Archive and can be accessed under the SRA accession number PRJNA393916.
B. mori
female and male HiSeq reads were produced on an Illumina HiSeq 2000 sequencer (101-bp pair-end using pools of larval posterior silk gland which yielded ∼73-fold coverage of genome for each sex).
B. mori
female MiSeq reads (only used for assisting
de novo
assembly of W contigs) were produced on an Illumina MiSeq sequencer (250-bp pair-end; pools of female larvae; ∼20-fold genome coverage). All reads were from Dazao, the inbred strain used in the genome project.
14
The next generation sequencing data for male and female
D. melanogaster
pooled 5 day old mated adults was downloaded from the NCBI Sequence Read Archive [SRA: SRP007888]; the next generation sequence data for male and female
An. gambiae
pooled wild type strain was downloaded from NCBI Sequence Read Archive [SRA: SRP014756].
12
,
15
Preferably, to ensure the accuracy of the KQ method, male and female sequence data should be from highly inbred populations and data for both sexes should be of the same amount. Since the coverage of the next-generation sequence data can affect the calculation of the KQ value, from our experience, we suggest that more than 50X coverage is an appropriate choice, which can produce a more precise high-frequency W-15-mer.
B. mori
RNA-Seq data of mixed sex embryos from 0–24 hpo were produced using an Illumina HiSeq 2000 sequencer, yielding 101-bp pair-end and 24 G clean data for each of the different periods (0, 2, 4, 8, 16 and 24 hpo). Small RNA sequences have been deposited in GenBank/EMBL/DDBJ under accession numbers from AB386191–AB424683.

The genome sequence of males and females differ only by the W chromosome, which produces W-specific
k
-mers. These k-mers are present only in the female data, but absent from the male data. Before doing a female
vs
. male
k
-mer comparison, removal of sequencing errors in the reads is important in order to achieve high resolution. This was done using two criteria: (i) masking low quality bases and (ii) filtering
k
-mers with rare frequency. Specifically, for the
B. mori
reads, the sequencing errors were filtered by masking bases with phred score <20 and removing 15-mers that were present fewer than 5X in reads. Counting
k
-mers for separate female and male data and removal of sequencing errors can be done using Jellyfish.
16
,
17
We set up the KQ parameter, for a given
k
-mer
K
i, KQ(
k
i) =
M
(
k
i)/
F
(
k
i), where
M
(
k
i) is the frequency of
K
i in male reads,
F
(
k
i) is the frequency of
K
i in female reads. To obtain reliable W-derived
k
-mers, we used the strict criterion of KQ = 0. To reduce the interference of individual polymorphism in sequencing reads, we filtered out these
k
-mers (KQ = 0) with low frequency (<15,
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) in female reads.

Alternatively, we can approach heaviness as an ideal, an essential quality or set of qualities that any kind of music can share to greater and lesser degrees. This is, I take it, what we hear in Sleep’s
Jerusalem
: the pinnacle of a musical genre that takes heaviness as its principal aesthetic ideal.
Jerusalem,
an album that Sleep explicitly intended to be “the heaviest thing ever recorded,” is a paradigm case of musical heaviness that challenges both the way most of us think and talk about heavy metal as well as the way that philosophers talk philosophically about rock music. In his 1993 essay, “
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,” for example, Bruce Baugh asks: “Can there be an aesthetics of rock music?” By this he means: Can there be an aesthetics of rock music as distinct from an aesthetics of classical music? His answer is yes, provided we can dispense with Kantian formalism and embrace the “
material
” (that is, the visceral, rather than intellectual) elements of rock music—
rhythm
,
notational expressivity
, and
loudness
—as the appropriate evaluative criteria.

Along these lines, we might think of heaviness as a
material
element of rock, something to be
felt
rather than cognized. But this view has its critics. For one, we need not think that visceral, non-formal features of music are exclusive to rock. (James Young
argues
, for example, that any of the features of rock music that seem to require a distinct aesthetics of rock music can also be found in classical music.) And second, the formal features of classical music can also be expressive qualities of rock. (Davies rightly
points out
that Baugh “underestimates the extent to which the visceral response he describes depends … on a song’s melodic and harmonic shape, its words, its overall structure, and so on,” in other words, its formal qualities.)

In contrast to this approach, it seems to me that heaviness is situated somewhere between the material and formal elements of music. What we find in Sleep’s
Jerusalem
is not a singular heavy-making property, but a combination of at least two properties:
sonic weight
and
sonic density
.

Sonic weight is materially similar to
rhythm
(not the formal, measurable quality of tempo but the perceptive quality of
timing
) in that its quality is literally felt in the body. Unlike rock music, however, the rhythm of doom metal does not inspire much movement or dancing. Clocking in at a lumbering 96 beats per minute (as compared with the average metal range of 250 to 500), the rhythm of
Jerusalem
is the perception of slowness. But heaviness is perceived not so much as a physical stimulation as an emotional or psychological weight pressing down on us—that is, as
sonic weight
. The music tends toward stasis rather than movement, and this resistance is amplified by the
simplicity
and
repetition
of the riff.

Sites 4, 13, 14, 18 and 19 contained individuals with mtDNA haplotypes from but the nuclear genome entirely from the other species. Jin . provided evidence that from the Qiadam Basin exhibited a rapid eastward range expansion as the Yellow River drained the paleao-Lake Gonghe, approximately 0.15 million years ago [
39
]. Under this scenario, would have come into secondary contact with populations isolated during this time. Hybridization would have resulted in introgression of mtDNA haplotypes from both of these species. Continual backcrossing with one parental species would have "diluted" any evidence of hybridization in their nuclear genomes, while the introgressed mitochondrial genome remained in place. Indeed, historical hybridization and introgression of mtDNA haplotypes has been reported in a number of studies ranging from amphibians to mammals [
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,
50
,
51
]. For example, Melo-Ferreira . reported extensive introgression in the Iberian Hares from Spain with 32% of containing an mtDNA lineage from [
49
].

Our genetic clustering did not match the morphological diagnosis provided by Wang . [
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], although the distribution of the genetic clusters was largely congruent with their proposed species ranges. Wang . suggested that has a tail length larger than its snout-vent length and has more than 100 rows of back scales, while has a relatively short tail and fewer than 100 rows of back scales [
35
]. Approximately half of identified in this study did not possess these "diagnostic" characters. This is not particularly surprising given that the genus is notorious for exhibiting large intraspecific morphological variation but lack of consistent interspecific differences [
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,
52
]. Such phenotypic diversity has made it particularly difficult to delineate species and morphology-based taxonomy is very confusing in this genus. For example, more than ten names have been applied to historically [
38
]. Rather than searching for a few diagnostic characters, a multivariate analysis, such as principle component analysis, may be more appropriate to dissect the morphological differences between and . It is important to note that a lack of congruence between morphology and other tools (e.g. genetic) for diagnosing species should not necessarily be regarded as evidence against the validity of two species [
8
]. Traditional morphology-based taxonomy, is only one way to describe "life's diversity" and species hypotheses developed based on morphology alone should be tested using different approaches with a variety of data sets [
8
]. Wang . also described chromosomal differences between the two species; while has ZW sex chromosomes, lacks identifiable sex chromosomes [
35
]. Unfortunately, we do not have chromosome data for specimens examined in this study. Nevertheless, the chromosome difference could potentially provide a reproductive isolation mechanism.